Ubiquitous eddies of the eastern Bering Sea and their coincidence with concentrations of larval pollock

Between 1988 and 1993, 12 satellite-tracked buoys were deployed in four eddies in the south-eastern Bering Sea. Our success in finding eddies resulted from placing buoys in high concentrations of walleye pollock (Them-gra chalcogramma) larvae. We utilize data from hydro-graphic surveys, satellite-tracked buoys and moored current meters to describe the eddies. Small (< 25 km diameter) eddies likely transit along the slope of the eastern Bering Sea every 45–60 days. In previous studies such small features were not observed because their size fell within typical separation of hydrographic stations and the weak sea surface temperature gradients are not resolved by satellite-borne infrared imagery.     

Ontogeny matters: Climate variability and effects on fish distribution in the eastern Bering Sea

Analyses of climate effects often ignore differences in life history for individual species. We analyzed a 34‐year time series of eastern Bering Sea fish surveys to evaluate changes in distribution by length and between cold and warm shelf‐wide average water temperatures for 20 species over inhabited depth, temperature, and location. All species showed evidence of ontogenetic migration. Differences in distribution between years with warm and years with cold shelf‐wide water temperatures varied among species and within species at different lengths. For species where shelf‐wide temperature effects were detected, the mid‐sized fish were most active in changing spatial distribution. For aquatic organisms ontogenetic migration occurs because life history stages have different environmental requirements. This study illustrates the need to consider species responses to climate change over different life history stages, and that studies on ecosystem responses should take ontogenetic differences into consideration when assessing impacts.     

Early- to late-summer population growth and prey consumption by age-0 pollock (Theragra chalcogramma), in two years of contrasting pollock abundance near the Pribilof Islands, Bering Sea

Acoustic survey data were used to estimate the abundance and distribution of age-0 walleye pollock and zooplankton near the Pribilof Islands, Bering Sea, nursery area at two time periods in two consecutive years: the beginning of August, and mid-September, of 1996 and 1997. The 1996 pollock year class ultimately produced a large adult cohort in the eastern Bering Sea, while the 1997 year class produced a below-average adult cohort. Acoustic densities of age-0 pollock were significantly lower in August – and declined more strongly from August to September – in 1997 than in 1996, indicating that the trend to adult cohort strength was already set by August. Diet composition analyses revealed that age-0 pollock ate a much higher proportion of euphausiids in 1997 than in 1996, despite lower acoustic abundance of euphausiids in 1997. We infer that in 1996, age-0 pollock experienced greater feeding success by August, with high concentrations of copepods available for smaller fish to consume, and high concentrations of euphausiids available for larger individuals. In 1997, age-0 pollock had lower body condition in August and may have been limited by the availability of small (<2 mm) copepods. Bioenergetic modeling of prey consumption did not indicate a likelihood that age-0 pollock would begin to deplete euphausiids until late August in 1996, and not at all between August and mid-September in 1997.     

Ichthyophonus‐infected walleye pollock Theragra chalcogramma (Pallas) in the eastern Bering Sea: a potential reservoir of infections in the North Pacific

In 2003, the Alaska walleye pollock industry reported product quality issues attributed to an unspecified parasite in fish muscle. Using molecular and histological methods, we identified the parasite in Bering Sea pollock as Ichthyophonus. Infected pollock were identified throughout the study area, and prevalence was greater in adults than in juveniles. This study not only provides the first documented report of Ichthyophonus in any fish species captured in the Bering Sea, but also reveals that the parasite has been present in this region for nearly 20 years and is not a recent introduction. Sequence analysis of 18S rDNA from Ichthyophonus in pollock revealed that consensus sequences were identical to published parasite sequences from Pacific herring and Yukon River Chinook salmon. Results from this study suggest potential for Ichthyophonus exposures from infected pollock via two trophic pathways; feeding on whole fish as prey and scavenging on industry‐discharged offal. Considering the notable Ichthyophonus levels in pollock, the low host specificity of the parasite and the role of this host as a central prey item in the Bering Sea, pollock likely serve as a key Ichthyophonus reservoir for other susceptible hosts in the North Pacific.     

A synthesis of biological and physical processes affecting the feeding environment of larval walleye pollock (Theragra chalcogramma) in the eastern Bering Sea

Biological and physical phenomena that affect conditions for larval survival and eventual recruitment differ in the oceanic and shelf regions. In the oceanic region, eddies are a common feature. While their genesis is not well known, eddies have unique biophysical characteristics and occur with such regularity that they likely affect larval survival. High concentrations of larval pollock often are associated with eddies. Some eddies are transported onto the shelf, thereby providing larvae to the Outer Shelf Domain. Advection, rather than local production, dominated the observed springtime increase in chlorophyll (often a correlate of larval food) in the oceanic region. Over two-thirds of the south-eastern shelf, eddies are absent and other phenomena are important. Sea ice is a feature of the shelf region: its interannual variability (time of arrival, persistence, and areal extent) affects developmental rate of larvae, timing of the phytoplankton bloom (and potentially the match/mismatch of larvae and prey), and abundance and distribution of juvenile pollock. In the oceanic region, interannual variation in food for first-feeding pollock larvae is determined by advection; in the shelf region, it is the coupled dynamics of the atmosphere–ice–ocean system.     

Anomalous conditions in the south-eastern Bering Sea, 1997: nutrients, phytoplankton and zooplankton

Anomalies in the regional weather over the south-eastern Bering Sea during spring and summer of 1997 resulted in significant differences in nutrient availability, phytoplankton species composition, and zooplankton abundance over the continental shelf as compared with measurements in the 1980s. Calm winds and the reduction of cloud cover in spring and summer produced a very shallow mixed layer in which nitrate and silicate were depleted after an April diatom bloom. High submarine light levels allowed subsequent phytoplankton growth below the pycnocline and eventual depletion of nitrate from the water column to depths of 70 m or more. Thus, total new production during 1997 may have exceeded that of previous years when nitrate was not depleted below the pycnocline. A bloom of the coccolithophorid, Emiliania huxleyi , was observed in early July in the warm, nutrient-depleted waters over the middle and inner shelf. Emiliania huxleyi concentrations reached 4.5 × 10⁶ cells L^–1 by September, and the bloom persisted through the autumn. There was evidence for increased abundance of some species of copepods in 1997 as compared with data from the middle domain in June 1981. The abundance of adult and juvenile euphausiids in 1997 was statistically similar to values measured in 1980 and 1981. However, near-surface swarms were rarely observed on the inner shelf in August–September 1997. Lack of euphausiid availability in the upper water column may partially explain the August–September mass mortality of planktivorous short-tailed shearwaters ( Puffinus tenuirostris ) observed on the inner shelf.     

Variation in the distribution of yellowfin sole Limanda aspera larvae in warm and cold years in the eastern Bering Sea

Multiyear periods of relatively cold temperatures (2007–2013) and warm temperatures (2001–2005 and 2014–2018) altered the eastern Bering Sea ecosystem, affecting ocean currents and wind patterns, plankton community, and spatial distribution of fishes. Yellowfin sole Limanda aspera larvae were collected from the inner domain (≤50 m depth) of the eastern Bering Sea among four warm years (2002, 2004, 2005, 2016), an average year (2006), and three cold years (2007, 2010, 2012). Spatial distribution and density of larvae among those years was analyzed using generalized additive models that included timing of sea-ice retreat, areal coverage of water ≤0°C, and water temperature as covariates. Analyses indicated a combination of temperature effects on the location and timing of spawning, and on egg and larval survival, may explain the variation in larval density and distribution among years. During warm years, higher density and wider spatial distribution of larvae may be due to earlier spawning, an expansion of the spawning area, and higher egg and larvae survival due to favorable temperatures. Larval distribution contracted shoreward, and density was lower during cold conditions and was likely due to fish spawning closer to shore to remain in preferred temperatures, later spawning, and increased mortality. Predicted drift trajectories from spawning areas showed that larvae would reach nursery grounds in most years. Years when the drift period was longer than the pelagic phase of the larvae occurred during both warm and cold conditions indicating that settlement outside of nursery areas could happen during either temperature condition.     

Copepod dynamics across warm and cold periods in the eastern Bering Sea: Implications for walleye pollock (Gadus chalcogrammus) and the Oscillating Control Hypothesis

Differences in zooplankton populations in relation to climate have been explored extensively on the southeastern Bering Sea shelf, specifically in relation to recruitment of the commercially important species walleye pollock (Gadus chalcogrammus). We addressed two research questions in this study: (i) Does the relative abundance of individual copepod species life history stages differ across warm and cold periods and (ii) Do estimated secondary production rates for copepods differ across warm and cold periods? For most copepod species, warmer conditions resulted in increased abundances in May, the opposite was observed in colder conditions. Abundances of smaller‐sized copepod species did not differ significantly between the warm and cold periods, whereas abundances of larger‐sized Calanus spp. increased during the cold period during July and September. Estimated secondary production rates in the warm period were highest in May for smaller‐sized copepods; production in the cold period was dominated by the larger‐sized Calanus spp. in July and September. We hypothesize that these observed patterns are a function of temperature‐driven changes in phenology combined with shifts in size‐based trophic relationships with primary producers. Based on this hypothesis, we present a conceptual model that builds upon the Oscillating Control Hypothesis to explain how variability in copepod production links to pollock variability. Specifically, fluctuations in spring sea‐ice drive regime‐dependent copepod production over the southeastern Bering Sea, but greatest impacts to upper trophic levels are driven by cascading July/September differences in copepod production.     

Spatial and temporal patterns in summer ichthyoplankton assemblages on the eastern Bering Sea shelf 1996–2007

Larval and early juvenile fishes were sampled from the eastern Bering Sea (EBS) shelf from 2001 to 2005, and in 2007. Data from these collections were used to examine spatial and temporal patterns in species assemblage structure and abundance. The years 2001–2005 were unusual because the EBS water temperature was ‘warm’ compared with the long‐term mean temperature. In contrast, 2007 was a ‘cold’ year. The abundance of the five most numerous taxa at 12 stations common to all years sampled (1996–2005, 2007) were significantly different among years. Larval and early juvenile stage Theragra chalcogramma (walleye pollock), a commercially important gadid, were by far the most abundant fish in all years. Bottom depth alone best explained assemblage structure in most years, but in others, bottom depth and water column temperature combined and percent sea‐ice coverage were most important. Abundance of T. chalcogramma larvae increases with water column temperature until 5°C and then becomes level. Higher abundances of Gadus macrocephalus (Pacific cod) larvae occur in years with the greatest percent sea‐ice cover as indicated by GAM analysis. Larvae of Lepidopsetta polyxystra (northern rock sole) increase in abundance with increasing maximum wind speed, but decrease at a later date during the last winter storm. The data are consistent with the hypothesis that oceanographic conditions, specifically water temperature and sea‐ice coverage, affect the spatial and temporal pattern of larval abundances. In general, ichthyoplankton species assemblages can be important early indicators of environmental change in the Bering Sea and potentially other subarctic seas as well.     

Multispecies biomass dynamics models reveal effects of ocean temperature on predation of juvenile pollock in the eastern Bering Sea

Walleye pollock (Gadus chalcogrammus) supports one of the largest commercial fisheries in the world. Juvenile pollock are important forage fish in the eastern Bering Sea (EBS) ecosystem, often representing the largest fraction in the diets of major Bering Sea piscivores. Large variability in the EBS pollock stock biomass in recent years has been attributed primarily to fluctuations in recruitment. It has been hypothesized that predation rates on forage fishes increase when the cold pool (a body of cold water < 2°C) is extensive and covers much of the middle continental shelf, which tends to concentrate larger predatory fishes in the outer shelf and slope regions. In contrast, young pollock appear to tolerate colder temperatures than older fish and can stay in the cold pool, thereby reducing predation. We used a multispecies modeling approach to examine the effects of the cold pool size on predation of juvenile pollock. We found that predation on age‐1 pollock by age‐3+ pollock decreased, and predation on age‐1 and age‐2 pollock by arrowtooth flounder increased with increasing bottom temperature, which was used as a proxy for the cold pool size. These results suggest that the cold pool creates spatial separation between juvenile pollock and arrowtooth flounder, but not between adult and juvenile pollock. The model developed in this study could be used to examine the effects of other covariates on interspecific interactions, help explain observed changes in fish communities, and understand implications of climate change on ecosystems and their productivity.     

Evidence for a substantial increase in gelatinous zooplankton in the Bering Sea, with possible links to climate change

We examined quantitative catches of large medusae from summer bottom trawl surveys that sampled virtually the same grid station on the eastern Bering Sea shelf using the same methodology every year from 1979 to 1997. This series shows a gradual increase in biomass of medusae from 1979 to 1989, followed by a dramatic increase in the 1990s. The median biomass increased tenfold between the 1982–1989 and 1990–1997 periods. Most of this biomass was found within the Middle Shelf Domain (50 <  z  < 100 m). The greatest rate of increase occurred in the north-west portion of this domain. Whether this dramatic increase in biomass of gelatinous zooplankton has resulted from some anthropogenic perturbation of the Bering Sea environment or is a manifestation of natural ecosystem variability is unclear. However, several large-scale winter/spring atmospheric and oceanographic variables in the Bering Sea exhibited concomitant changes beginning around 1990, indicating that a possible regime change occurred at this time.     

Spatial and temporal patterns in summer ichthyoplankton assemblages on the eastern Bering Sea shelf 1996–2000

Larval and early juvenile fishes were sampled from the eastern Bering Sea (EBS) shelf during summer from 1996 to 2000. Data from these collections were used to examine spatial and temporal patterns in species assemblage structure and abundance. Cluster analyses based on Bray–Curtis dissimilarity coefficients were used to group species and stations according to similar abundance and species composition. Ordination techniques were used to verify groupings, and a non‐parametric stepwise procedure using a Spearman correlation coefficient (BIO‐ENV) was used to relate groupings to predominant environmental variables. These approaches revealed a pattern of station groupings that were generally related to bathymetry in 1996, 1997, 1999, and 2000, although no obvious relationship to geographic boundaries was observed in 1998. Significant differences in species associations were observed in 1997 and 1998, and depressions in abundance were also noted among many species between 1997 and 1999. A regional, full primitive equation model was used to simulate float trajectories on the EBS shelf in each year to better relate fish distributional observations to prevailing current patterns. Model results indicated general variations in flow in several years, although 1998 stood out with stronger northeast flow than in any of the other years examined. Observed disruptions of larval and early juvenile fish assemblages could be related to the strong El‐Niño event of 1997–98 in the EBS. If this idea is confirmed, our study suggests that larval and juvenile fish are sensitive and respond relatively quickly (1–2 yr) to environmental perturbations, and as such, may be timely indicators of environmental change.     

Predation on walleye pollock (Theragra chakograrnma) eggs and yolk-sac larvae by pelagic crustacean invertebrates in the western Gulf of Alaska

Immunological detection of yolk protein was used to assess predation by pelagic amphipods (gammarid and hyperiid), mysids, and euphausiids on eggs and yolk-sac larvae of walleye pollock Theragra chalcogramma during 1988 and 1989. Consumption estimates were made on the basis of frequency of positive immunoassays, assay detection times (gut clearance time), predator abundance, and spatial overlap of predators and prey. From our results gammarid amphipods and euphausiids were important predators on eggs and yolk-sac larvae, respectively. Gammarid amphipods alone consumed about 14% of the standing stock of pollock eggs in 1989. These results were compared with those from clearance rate experiments of predators feeding on pollock eggs in 300-1 bags. In general, clearance rate estimates of egg consumption were lower than those determined from gut contents.     

Interannual variability in growth of walleye pollock, Theragra chalcogramma, in the central Bering Sea

Interannual variability in growth of walleye pollock, Theragra chalcogramma, was examined. Adult walleye pollock were collected from the central Bering Sea (Aleutian Basin) from 1978 to 1999. Average fork lengths were found to be approximately 47 cm during the 1970–80s, this increased to 56 cm in the late 1990s. Age was determined for 4805 individuals using the otolith break and burn method. Ages ranged from 5–23 years and the year classes of 1978 and 1989 were dominant in the 1980s and the 1990s, respectively. Fish had significantly larger length-at-age in the 1990s compared to the 1970–80s, and interannual variability in age–length relationship was clearly observed. Taking into consideration a recent decrease of the walleye pollock biomass in the central Bering Sea, density-dependent growth was supported as one possibility of the growth variability. At the same time, we could not rule out the possibility that oceanographic variability affected the growth of walleye pollock in the area.     

Potential movement of fish and shellfish stocks from the sub‐Arctic to the Arctic Ocean

An assessment of the potential for 17 fish or shellfish stocks or stock groups to move from the sub‐Arctic areas into the Arctic Ocean was conducted. A panel of 34 experts was convened to assess the impact of climate change on the potential movement of the 17 stocks or stock groups. The panel considered the exposure of species to climate change, the sensitivity of species to these changes and the adaptive capacity of each stock or stock group. Based on expert opinions, the potential for expansion or movement into the Arctic was qualitatively ranked (low potential, potential, high potential). It is projected that the Arctic Ocean will become ice‐free during the summer season, and when this happens new areas will open up for plankton production, which may lead to new feeding areas for fish stocks. Five stocks had a low potential to move to, or expand in, the high Arctic. Six species are considered as potential candidate species to move to, or expand in, the high Arctic. Six stocks had a high potential of establishing viable resident populations in the region. These six stocks exhibit life history characteristics that allow them to survive challenging environmental conditions that will continue to prevail in the north. This study suggests that several life history factors should be considered when assessing the potentiality of a species moving in response to changing climate conditions.     

Anomalous conditions in the south-eastern Bering Sea 1997: linkages among climate, weather, ocean, and Biology

In 1997, the Bering Sea ecosystem, a productive, high-latitude marginal sea, demonstrated that it responds on very short time scales to atmospheric anomalies. That year, a combination of atmospheric mechanisms produced notable summer weather anomalies over the eastern Bering Sea. Calm winds, clear skies, and warm air temperatures resulted in a larger-than-normal transfer of heat to surface waters and the establishment of a shallow mixed layer. In spring, significant new production occurred below the shallow pycnocline over the Middle Shelf, depleting the subpycnocline nutrient reservoir that normally exists during summer. Following the depletion of nitrate and silicate from the system, a sustained (≥ 4 months) bloom of coccolithophores ( Emiliania huxleyi ) was observed – a phenomenon not previously documented in this region. Summer Middle Shelf Domain copepod concentrations were higher for some species in 1997 than in the early 1980s. Warmer surface water and lack of wind mixing also changed the basic distribution of hydrographic regimes on the south-eastern shelf and altered the strength and position of fronts or transition zones where apex predators seek elevated food concentrations. The Inner Front was well inshore of its normal position, and adult euphausiids (the primary prey of short-tailed shearwaters, Puffinus tenuirostris ) were unavailable at, and shoreward of, the front in autumn. High shearwater mortality rates followed the period of low euphausiid availability. Some, but not all, of these anomalous conditions re-occurred in 1998. These observations are another demonstration that the structure and function of marine ecosystems are intimately tied to forcing from the atmosphere. Alteration of climatological forcing functions, expressed as weather, can be expected to have large impacts on this ecosystem and its natural resources.     

Causes of walleye pollock (Theragra chalcogramma) recruitment decline in the northern Sea of Japan: implications for stock management

Recruitment of the northern Japan Sea stock (JSS) of walleye pollock has been decreasing since around 1990. In this study, I analyzed the factors causing this decrease in recruitment by investigating the relationship between recruitment, spawning stock biomass (SSB) and environmental factors using a generalized additive model (GAM). GAM fit to the data showed the importance of SSB, sea surface temperature (SST), ocean current strength (Tsushima Warm Current) and wind intensity (Asian monsoon) in determining the recruitment. Of these, the relationship between SSB and recruitment was positive and not negatively density‐dependent. On the other hand, the recruitment was negatively related to SST and ocean current strength, and a dome‐shaped relationship was observed between wind intensity and recruitment. Since around 1990, the values of SST and ocean current strength have mostly been high and that of wind intensity mostly low. In addition, SSB has been decreasing since the late 1990s. It is likely that the recruitment decline of JSS after approximately 1990 has been caused by warm water temperature, strong Tsushima Warm Current and weak Asian monsoon, and that the recent decrease in SSB has amplified this recruitment decline. According to the model’s estimation, a recruitment recovery due to environmental improvement will be highly restricted as long as SSB remains at its current low level. Significant recovery of SSB is urgently needed for JSS.     

A review: Walleye pollock in the North Pacific–how much difference do they really make?

Populations of several species of marine birds and mammals in the Bering Sea and Gulf of Alaska have been declining since the mid-1970s, with numbers of one, the Steller sea lion (Eumetopias jubatus) , so depressed it was listed as threatened under the Endangered Species Act in spring 1990. All of the declining populations depend to an important extent on walleye pollock (Theragra chakogramma) for food, although they eat numerous other species as well. In contrast, certain animals that compete with pollock for common prey have been increasing in abundance. All of these changes could be related through food web connections mediated by pollock. Pollock is also important to people–it presently supports the largest single-species commercial fishery in the world, in large part because of its great biomass, which has averaged about 15 × 10⁶t in the Bering Sea over the past 15 years. Pollock consume an inordinate proportion of the pelagic production in the Bering Sea, which further supports the conclusion that it is a key species in the ecosystem. However, there are conflicting hypotheses about the importance of the roles played by pollock as predator and prey, and about the effect that changes in pollock abundance might have on biomass yield at higher trophic levels.     

On the variable subsurface environment of fish stocks in the Bering Sea

Data from 1971 to 1993 are used to examine characteristics and variability of the subsurface temperature maximum in the Aleutian Basin of the Bering Sea. The temperature of this feature varies from ∼3.5C to 4.6C at depths of ∼150–400 m, and sigma-t density is usually in the range 26.6–26.9. The variability is caused mainly by the absence or presence of inflow of warm, low-density Alaskan Stream water through Amukta Pass. Thus the environment offish stocks is far from uniform.